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 Fig. 318.—A Diagram of an Orchid-flower.
B, Cephalanthera. Stylar-column: a anther; s
stigma; at the foot are seen scars indicating the
position of the parts which have been removed.
Order 2. Orchidaceæ. The epigynous, petaloid perianth is
strongly zygomorphic in having the posterior leaf of the interior whorl,
the lip (labellum), differing from all the other leaves in form, size, and
colour (except Apostasieæ); the position of the labellum is very
frequently reversed, being turned forwards and downwards by the
twisting of the ovary (Fig. 318 A). Only 1 of the stamens—the
anterior of the external whorl—is developed and bears an anther (by
the twisting of the ovary it is turned posteriorly and upwards); the
others are entirely wanting (indicated by * in Fig. 318 A) or present
as staminodes (Fig. 318 A, δ δ) (except Apostasieæ, Cypripedileæ);
the filaments are united with the style to form a column (Fig. 318 B),
the stylar-column[30] (gynostemium), and the anther (a) is thus
placed on its apex and exactly behind or over the stigma (s). The
anther is 4-locular; the pollen-grains do not separate (except
Apostasieæ, Cypripedileæ) but remain united either in tetrads or in
masses, which correspond to a pollen-mother-cell (Fig. 320 C, D, E);
or the pollen-grains, formed in each of the two anther-halves, remain
united and form one or a few wax-like masses (pollen-masses,
pollinia). The 3 carpels form a unilocular ovary with 3 parietal, deeply
bifid placentæ (except Apostasieæ, Selenipedilum). Only the two
lateral carpels are prolonged and developed into the stigma (Fig. 318
B, s), while the one lying in the median line, which is situated just
within the anther (Fig. 318 A), becomes either rudimentary or
developed into the “rostellum” (“a small beak”), on which the sticky
bodies (glandulæ) arise; by aid of these the heavy, connected pollen-
masses may be glued to the insects which visit the flower, and
pollination is thus secured (in Apostasieæ and Cypripedileæ the 3
carpels each contribute to the formation of the stigma). The fruit is a
capsule which most often dehisces by 6 valves, 3 of which are
broader and bear the placentæ, and 3 alternating with them are
narrower and barren (except Vanilla). The very numerous and
exceedingly small seeds have no endosperm, and have a somewhat
spherical embryo without any trace of external organs. The testa is
membranous and loose.
The Orchids are all perennial herbs with diverse habits and
varying morphological structure (see the genera); the leaves are
scattered, of the usual Liliaceous form, and the inflorescences in all
cases are racemes or spikes (sometimes branched), with subtending
bracts, but without bracteoles.
The forms which are the least modified are described first.
1. Apostasieæ. The perianth-leaves are almost alike and free.
The column is straight, with 3 equally-developed stigmas. Neuwiedia
has 3 perfect stamens (1 median of the outer whorl, and 2 lateral of
the inner whorl); Apostasia has only 2 perfect (inner lateral) and one
barren (the median of the outer whorl), which however may be
entirely wanting. The 3 posterior stamens are entirely suppressed.
The pollen is powdery. The ovary is 3-locular with axile placenta. 7
species (Tropical East India, Australia).
 Fig. 319.—Cypripedilum calceolus: 1 front view of the flower; 2
lateral view, after the removal of all the perianth-leaves with the
exception of the labellum, which has been divided longitudinally; 3
the stylar-column; ov ovary; s-s exterior, p interior perianth; p’ the
labellum; a the two fertile stamens; a’ the staminode; st the stigma;
i entrance for the insects; ex exit.

2. Cypripedileæ.[31] The flower is strongly zygomorphic with a

large boat-shaped labellum. There are two perfect stamens
belonging to the inner whorl, and the median anterior (later on the
posterior) stamen of the outer whorl is transformed into a large,
barren, shield-shaped body (Fig. 319). Selenipedilum has a 3-locular
ovary, but Cypripedilum (Ladies’-slipper) has a unilocular ovary with
3 parietal placentæ—the typical structure for the Orchids. The
pollen-grains are separate (not in tetrads) and all the 3 lobes of the
stigma are constructed to receive them. This group is therefore, next
to the Apostasieæ, the least modified among the Orchids; in all the
following groups, one of the lobes of the stigma is differently
developed from the others, and there is only one stamen.—Terrestial
Orchids.—The pollination of C. calceolus is effected by the forcible entrance of
insects into the boat-shaped labellum (Fig. 319 p’) at i, and their escape at ex (in
2) where the anthers are situated; in this way the stigmas will first be touched and
then the anthers. The pollen-grains are surrounded by a sticky mass in order that
they may adhere to the insects.
3. Neottieæ. The majority are terrestrial Orchids with creeping,
sympodial rhizomes; the blades of the leaves are not detached from
the stem at joints, and have convolute vernation. The anthers do not
drop off, but persist in the withered condition; their apex is brought in
contact with the rostellum (acrotonous Orchids). The pollen-grains
are united in tetrads, which, however, often hang loosely together in
pollinia, attached to a sticky part of the rostellum (“adhesive disc”),
so that they adhere to the insects, and are by them transferred to the
stigmas. Spiranthes. Listera; Neottia. N. nidus-avis (Bird’s-nest) is brown (it
has little chlorophyll) in colour, has no foliage-leaves, and lives mainly as a
saprophyte; the rhizome is studded with unbranched, fleshy roots which may form
buds at their extremities.—Vanilla climbs by aerial roots. The fruit is
fleshy and hardly opens, or does so irregularly.—Epipactis,
Cephalanthera.—Epipogon and Limodorum are saprophytes without
chlorophyll.
 Fig. 320.—A Flower of Orchis maculata (front view): a stamen; b the cup; n the
stigmas; x staminodes; sp the spur; spe the entrance to it; sm-sl-sl exterior
perianth-leaves; pm the labellum, and pl-pl the other 2 interior perianth-leaves. B-E
Orchis mascula: B lateral view of the column; C a pollinium with massulæ (p),
caudicle (c) and adhesive disc (d); D caudicles with the cup (r), front view; the
latter is depressed so that the adhesive disc is seen lying inside it; E a pollinium,
more highly magnified; some massulæ are removed. F Ophrys aranifera: rostellum
and the base of the anther-loculus; an adhesive disc is seen on the right.
4. Ophrydeæ. Anthers 2-locular, not falling off, on a very short
column. The anther is united at its base with the rostellum
(basitonous Orchids, Fig. 320 A, B), while in all other Orchids it is
connected at the apex (acrotonous Orchids). The pollen-grains in
each loculus are united into small “masses” (massulæ), each of
which corresponds to a pollen-mother-cell in the anther, and which
hang together by elastic threads (Fig. 320 C, E). Each pollinium is
attached at the base by a stalk (caudicle) to an adhesive disc,
formed by the modified stigma (rostellum), and is easily liberated
from it (Fig. 320 C, D, F). The pollinium, which is formed in an
anther-loculus, together with its caudicle and adhesive disc, is
termed “pollinarium” (Fig. 320 C).—All Ophrydeæ are terrestrial with
tuberous roots, two of which are present in the flowering period, an
older one (from the preceding year) containing the nourishment for
the flowering-shoot of the year, and a young one which is intended to
contain the reserve material for the following year. Inflorescence
terminal.
Orchis. The lip has a spur; each of the club-like pollinia is
attached to its own adhesive disc, the discs being enclosed in a
common pouch formed by the rostellum (Fig. 320 C, D). Tubers ovate,
undivided: O. morio, mascula; tubers palmate: O. incarnata, maculata, majalis.—
Ophrys; no spur, the two adhesive discs are each enclosed in a
separate pouch (Fig. 320 F).—Anacamptis and Serapias have one
adhesive disc.—Habenaria, Gymnadenia, Platanthera, Herminium,
Nigritella, Cœloglossum, etc., have naked adhesive discs (no
rostellum).
5. Epidendreæ. Acrotonous Orchids with deciduous anthers (except Malaxis);
2-8 wax-like pollinia, with or without caudicles; generally no adhesive discs.
Malaxis (the flower is twisted through a complete circle, causing the labellum to be
turned upwards), Sturmia and Corallorhiza[32] (Coral-root); the latter has a
creeping, coral-like rhizome without roots, and is destitute of chlorophyll except in
the ovary. The other two somewhat resemble the tropical Orchids in having the
lower internodes of the axis of the inflorescence tuberous. Liparis; Calypso. Most
of the genera are tropical epiphytes and many have aerial, green tubers formed
from one or more stem-internodes; Dendrobium, Eria, Phaius, Bletia, Epidendrum,
Cattleya, Lælia, Pleurothallis, Restrepia, Masdevallia, Bulbophyllum, etc.
6. Vandeæ. These resemble the preceding but have only 2 wax-like pollinia in
each anther, which are attached by a caudicle to the adhesive disc of the
rostellum. Nearly all are tropical epiphytes. Stanhopea, Catasetum, Maxillaria,
Oncidium, Vanda, Polystachya, etc.
6,000 (10,000?) species. The majority live in the Tropics and occur, especially,
as epiphytes on trees or in the crevices of rocks, to which they are attached by
aerial roots. These aerial roots, like those of Araceæ, are covered by several
layers of spirally-thickened cells (tracheides) which contain air and form the
velamen—an apparatus to absorb moisture from the air. The roots have a white
appearance when the cells are filled with air, which changes to a greenish hue
when they are filled with water, the chlorophyll then shining through. They
generally have horizontal rhizomes; the ascending shoots, which bear the foliage-
leaves, may vary, but they very often swell and assume the form of a tuber, which
persists for several years fresh and green after the leaves have fallen off (Fig.
321). Vanilla is an exception (see above). Our Orchids are all terrestrial (or marsh-
plants); the largest number of species is found in calcareous soils.
 Pollination takes place principally by means of insects, but self-pollination
occurs in some. The lip serves as a landing-stage for the insect visitors, which, on
sucking the honey, cause the adhesive discs, with the pollinia attached to them, to
adhere to their bodies (generally to the probosces) and so carry them away to
other flowers. In some species parts of the flower are sensitive or irritable, which
has some connection with the pollination. Without doubt there are a great many
biological differences which are closely connected with the infinite multiplicity of
forms; Darwin (1862) has already shown an enormous variety, never even dreamt
of before, in the European species. The genus Catasetum has ♂-♀-and ☿-plants
with flowers of such different appearances that they have been classed in various
genera (Myanthus, Monacanthus). Platanthera is pollinated by hawk-moths;
Ophrys, by flies; Epipactis latifolia, by wasps; Orchis, by bees, especially humble-
bees, etc.

Fig. 321.—Chysis bractescens.

The distribution of seeds is effected by the wind, the seeds being so
exceedingly small and light. Many species moreover have peculiar, elater-like, fine,
hygroscopic hairs in the ovary, which eject the seeds in a manner similar to the
elaters of the Liverworts.
The uses are few, mostly as ornamental plants in conservatories. The tubers of
several Orchis-species are officinal; they contain starch and mucilage and are
used us “salep.” The fruits of Vanilla planifolia are used as condiments and differ
from other Orchid-fruits in being rather fleshy and in dehiscing irregularly; the
seeds are very small, shining and black.

Class II. Dicotyledones.

In this class the embryo has 2 seed-leaves, a rule from which
there are few exceptions (e.g. Ficaria, Cyclamen, Pinguicula, certain
species of Corydalis, with only 1; and a few, mostly parasitic forms,
e.g. Monotropa, Orobanche, Pyrola, entirely without cotyledons). On
germination the cotyledons nearly always raise themselves above
the ground as green, assimilating leaves and are then termed aerial
or epigean, in contradistinction to the underground or hypogean
which are always buried. The structure of the seed varies
(endospermous or exendospermous); the embryo may be straight or
curved. In many instances the primary root grows as a vigorous tap-
root, with weaker branches arising acropetally (in annuals, biennials,
many perennials, especially woody plants); but in a large number of
herbaceous perennials, which have rhizomes, the root behaves very
much as in the Monocotyledons. The roots generally increase in
thickness by means of a cambium.
The stem, when seen in transverse section, has its vascular
bundles arranged in a ring; in reality, however, they form a kind of
cylindrical network in the stem; the bundles are open, and thickening
takes place by means of a cambium; annual rings are formed in the
perennial stems. There is a rich and very varied form of branching.
The two first leaves of a shoot (fore-leaves) are placed nearly always
to the right and to the left; the same arrangement is found in the two
first leaves developed on the flower-stalk, and these are, as a rule,
the only two; they are found below the calyx and are usually termed
the “bracteoles.” It has become customary to indicate the bracteoles
by the letters α and β, according to their sequence of growth, and in
that sense these letters will be employed in the following diagrams.
The arrangement of the leaves varies very much; there is also
a great variety of shapes in the leaves and their venation, but the
linear leaves, with parallel venation, so frequent in the
Monocotyledons, are seldom met with, as also the large sheaths
(though the sheath is well developed in the Umbelliferous plants);
stipules occur much more frequently.
The flower is most commonly cyclic, but acyclic or hemicyclic
forms also occur. The type which may be taken as a basis consists
in the majority of instances, as in the Monocotyledons, of 5 whorls, of
which the 4 outer ones (calyx, corolla, and the 2 whorls of stamens)
are most frequently 4 or 5 in number and placed in regular
alternation, whilst the innermost one (the carpels) has generally
fewer members, probably on account of space (Figs. 360, 361, 421,
429, 487, etc.). Trimerous (Figs. 384, 387, etc.) flowers, or those in
which the members of the flower are in threes or a multiple of three,
also occur, as well as dimerous flowers; other numbers are rare. It is
of the greatest importance in connection with the relative position of
the members of the flower to the axis and bract (orientation),
whether the bracteoles are typically present (even though they may
not be developed), or are typically absent. If there are 2 bracteoles
present, then their position in a pentamerous flower is often as
follows: the first sepal turns obliquely forward, the second is posterior
and median, the third obliquely forward, the fourth and fifth obliquely
backward; quincuncial æstivation is often found in these buds (Figs.
360, 429, 471, 475, 584). The first and third leaves, in the following
chapters, are most frequently alluded to as the “anterior,” the fourth
and fifth as the “lateral” leaves. The reversed arrangement, with the
median sepal in the front, occurs for instance in Papilionaceæ (Fig.
511), Lobeliaceæ (Fig. 594), Rhodoracecæ. If any bracteoles are
present below a tetramerous flower, the relation is generally that 2
sepals (the first ones) stand in the median plane, the two next ones
transversely (Fig. 393), and the corolla then adopts a diagonal
position (Fig. 397); but a diagonal position of the calyx generally
shows that the flower is not, strictly speaking, tetramerous, as in
Plantago (Fig. 567), Veronica (Fig. 559 C) and others.
If the bracteoles are not typically present, then the position of the
sepals is changed accordingly, and the two outer sepals endeavour
to assume the position which the bracteoles would otherwise have
occupied, e.g. in Primula (Fig. 547). Other positions are also found
when the number of bracteoles is more or less than two.
 The leaves which follow the sepals occupy definite positions with
regard to them, which we may consider later. An arrangement must,
however, be mentioned here; when the flower is “diplostemonous”
that is, has two whorls of stamens (thus, Sn, Pn, An + n), these may
be arranged in two ways. Either the first-formed whorl of stamens,
which are termed the “calyx-stamens,” stands directly in front of the
sepals (that is “episepalous”), and is the outermost whorl, and in this
case a regular alternation takes place between sepals, petals and
the two whorls of stamens, which is also continued into the carpels if
their number is the same as that of the other whorls: the carpels are
then placed opposite the sepals (Fig. 278) and the flower is
isomerous and Gn should be added to the formula above. Or, the
calyx-stamens form the innermost whorl, and the corolla-stamens,
which are subsequently formed (“epipetalous” stamens), stand
outside these (Figs. 360, 429); if the number of carpels is the same
as that of the preceding whorls, they are often placed right in front of
the petals and the corolla-stamens. The first-mentioned arrangement
is termed Diplostemonous, and the second Obdiplostemonous. Both
arrangements may be found in one and the same order, e.g. Caryophyllaceæ. The
size and relation of the members of the flowers, and also the contact with other
members in the early stages of their development, play an important part in
determining the arrangement.
The great number of structural arrangements found in this
enormously large class, may, as is the case in the Monocotyledons,
be further varied by suppression and division of certain leaves
(especially the stamens). Instances of this will occur in the following
(Figs. 559, 568.—426, 441, 445, etc.).
The Dicotyledons were formerly divided into 3 sub-classes:
Apetalæ (those without corolla), Sympetalæ or Gamopetalæ (those
with the petals united), and Choripetalæ or Polypetalæ (the petals
not united). This division has now been abandoned because it has
been proved that the Apetalæ were merely reduced or incomplete
forms of the Choripetalæ, and they have therefore been distributed
among the various families of the latter sub-class.
With regard to the Sympetalæ (or Gamopetalæ) it may be stated
that they form to a very great extent a closely connected and natural
group, having in common not only the character that the corolla is
gamopetalous and the stamens united with it (this being also found
in the Choripetalæ), but also a great many others (such as persistent
calyx, cyclic flowers with the formula S5, P5, A5 and as a rule G2,
the two carpels being united to form the ovary; seeds with a thick
integument and a very small nucellus). They are therefore
considered as an independent sub-class, and must be placed at the
close of the system of classification as the forms which presumably
have arisen the latest. In the future systems of classification this
arrangement will very probably be changed, and the first families of
the Sympetalæ, the Bicornes and others will for instance be to a
certain extent united with the families or orders of the Choripetalæ.
The Sympetalæ may certainly be considered as the youngest types,
the strongly pronounced metamorphosis supporting this theory, as
also the formation of the integument of the ovule, the one thick
integument being undoubtedly derived from the coalescence of two
—a holochlamydeous ovule, etc.
The Apetalæ and Choripetalæ are united into one sub-class. The
leaves of the perianth in this case are, as a rule, free from each
other, the structure of the flowers presents many differences, and the
ovules have as a rule 2 integuments and a large nucellus.
Considerable uncertainty still prevails regarding the arrangement
and the relationship of the individual families of the Choripetalæ, and
some of the following families are hardly quite natural; but the best
arrangement arrived at so far has been adopted here.
At the beginning of the book a review of the orders of the
Dicotyledons will be found.
Sub-Class 1. Choripetalæ. Petals free.

Family 1. Salicifloræ.
Trees and shrubs, which, in the structure of the vegetative shoot
and the catkin-like inflorescences, resemble the Quercifloræ, but the
structure of the flower differs so much from them, that the only order
brought under this heading—Salicaceæ—well deserves to be
separated and to form a family of its own, the nearest relatives of
which are still doubtful. As Juglandaceæ and Myricaceæ also deserve to be
placed in a special family, the name Amentaceæ (Catkin-bearers), hitherto applied
to all of these plants, cannot be retained as the name of a family.

Fig. 322.—Male and female catkins of Salix caprea.

There is only one order.
Order. Salicaceæ (Willows). Trees with simple, scattered,
stipulate leaves. Diœcious. The flowers are arranged in simple
inflorescences (spikes or racemes) which are termed catkins, and
which fall off as a whole after flowering (♂) or after the ripening of the
fruit ( ♀ ) (Fig. 322). The perianth is very imperfect[33] or wanting,
particularly in Salix (Fig. 323 o); the ♂-flower with 2–several stamens
and without any trace of a carpel (a, b, c): the ♀ -flower has a free
bicarpellate ovary, unilocular, and formed from 2 lateral carpels with
2 parietal (median) placentæ and generally ∞ ovules; the style
divides into two stigmas (d, e, f). The fruit is a two-valved capsule
and the very small seeds bear a tuft of hairs at the base. Endosperm
absent.—The catkins are situated on dwarf-branches, which in some species
often develop before the leaves and bear at their base only scale-leaves; in others
foliage-leaves are borne beneath the catkins. The vegetative bud commences with
2 bud-scales which are united on the anterior side into a scale. The capsule opens
by the dorsal suture. The seed-hairs spring from the funicle.

Fig. 323.—Salix: male flowers of S. pentandra (a), S. aurita (b), S. rubra (c),
female flowers of S. aurita (d), S. nigricans (e), S. mollissima (f).
Salix (Willow) has short-stalked, most frequently lanceolate leaves
and erect catkins with undivided bracts (Fig. 322). The flowers are
naked; 1 (o in a-f) or 2 yellowish glands situated in the median line.
In the ♂ -flower generally two stamens, situated laterally like the
carpels in the ♀-flower. Various forms are seen in Fig. 323.—The terminal bud
of the branches often aborts regularly, the uppermost lateral bud taking its place.
Populus (Aspen, Poplar) has long-stalked, more or less round or
cordate leaves with drawn-out apex; catkin pendulous; lobed bracts;
perianth cup-like with oblique edge; stamens usually numerous;
stigmas often divided.—P. tremula (Aspen) has received its name from the
tremor of the leaves: cf. “to shake like an aspen leaf.”
Pollination. The Poplars are wind-pollinated. The Willows have sticky pollen
and are pollinated by insects. The catkins of the Willows, especially the ♂ , are
more conspicuous, from the numerous, closely-packed, yellow flowers, rich in
honey and pollen. The catkins often appear before the foliage and so are much
more easily seen, whilst at this time of the year the number of competing honey-
flowers is smaller, and the insect visits consequently more numerous. On many
catkins of the Willow the flowers open earliest on the side which is turned towards
the sun and in descending order, i.e. the upper flowers develop before the lower
ones. Hybrids frequently appear.
 There are about 180 species existing in the northern, cold and temperate
latitudes. Some in the Polar regions are scarcely more than an inch in height, and
have a creeping rhizome (Salix herbacea, polaris, reticulata). Fossil forms are
found in the Tertiary and perhaps also in the Upper Cretaceous.
Uses. Principally for ornamental trees, as they grow very quickly and are easily
propagated by cuttings, S. babylonica, Weeping Willow; S. purpurea; Populus
alba, Silver Poplar; P. pyramidalis, Pyramid Poplar—a form of P. nigra; P.
monilifera, Canadian Poplar. The wood is very poor and little used; the branches of
many Willows are cultivated for basket-making, etc. The wood of the Aspen is
used for matches. The bark contains tannin and, in many Willows, a very bitter
extract, Salicin (S. pentandra, fragilis). Salicylic acid (officinal) is obtained from
Salix. Balsam is extracted from the buds of many Poplars, especially when the
leaves are shooting.

Family 2. Casuarinifloræ.
Trees with verticillate, scale-like leaves forming sheaths at the
nodes. Monœcious. Flowers unisexual. ♂ -flowers in catkins; ♀ in
short spikes. Pollen-tube entering the ovule at the chalaza, and not
through the micropyle. Ovary 1-seeded, unilocular. Carpels uniting
into a multiple fruit. Only one order.
Order. Casuarinaceæ. Trees (30 species), from Australia and
certain parts of S.E. Asia, with peculiar, equisetum-like appearance.
The leaves are verticillate, scale-like and united into sheaths. The
internodes are furrowed. Branching verticillate. The unisexual
flowers are situated in catkins or short spikes. The ♂ -flower has a
central stamen, surrounded by 2 median, scale-like perianth-leaves
and 2 lateral bracteoles. The ♀-flower has a 1-chambered ovary (2
ascending, orthotropous ovules), no perianth, but 2 large, lateral
bracteoles which finally become woody and form two valves,
between which the nut-like fruit is situated. The multiple-fruits
therefore resemble small cones.—Casuarina equisetifolia, cultivated,
gives “iron-wood.”
[The Casuarinas differ from the ordinary Dicotyledons in many important
respects which may be briefly summarised thus:—The bicarpellate ♀-flower has a
well-pronounced stylar-cylinder terminated by two stigmas, but the cavity of the
ovary closes very soon after its formation, and in it are developed two parietal
ovules; these are united by a bridge of cellulose to the stylar-cylinder or summit of
the ovary, and hence the ovules are connected with the walls of the ovary by the
bridge (above), as well as by the funicle (below). The archespore is developed
from the hypodermal cells at the summit of the nucellus, two primordial mother-
cells are first formed and from these by tangential divisions a central cylindrical
mass of cells (sporogenous-tissue) is produced which is surrounded by tapetal
cells. The cells of the sporogenous tissue correspond to the mother-cells of the
embryo-sac of other Angiosperms; they divide transversely and from 16–20
macrospores are formed together with inactive cells which are not crushed
together as in the case of other Phanerogams. The sexual apparatus is developed
from a single cell, but the number of cells composing this apparatus is subject to
variation, the oosphere being accompanied by one or two neighbouring cells which
resemble canal-cells rather than synergidæ. The sexual apparatus is found in the
majority of the macrospores, but in most of these it remains as a number of naked
cells; while in the fertile macrospores the cells are invested by walls of cellulose
(usually only one fertile macrospore is found in each ovule). Antipodal cells are
never developed. The macrospores elongate considerably towards the chalaza,
into which some penetrate. The pollen-tube traverses the stylar-cylinder and
enters the ovules at the chalaza, its passage through the tissue of the nucellus
being assisted by the prolongation of the macrospores. About the centre of the
nucellus the pollen-tube is ruptured; the apical portion which alone takes part in
the fertilisation being firmly attached to the macrospore. Although the actual
impregnation has not been observed, Treub considers that the endosperm begins
to be formed before fertilisation.]

Family 3. Quercifloræ.
Trees and shrubs with small, unisexual, monœcious flowers,
having no perianth or a simple inconspicuous one. The ♂ and ♀
flowers are very different and generally placed in separate
inflorescences. The ♂ -flowers are most often adnate to the bracts.
The stamens are placed opposite the perianth-leaves, when they are
present in equal numbers. The ♀-flower is naked, or has a superior
perianth. The ovary at the base is 2- or 3-(-6) locular with 1 or 2
pendulous ovules in each loculus, only one of which is developed;
the fruit is a one-seeded nut; endosperm absent; embryo straight.
The inflorescences, which are either compound and mixed (small
dichasia in spikes) or simple, are here also termed catkins; but,
strictly speaking, this term is applied to the ♂-inflorescences only. In
all Quercifloræ the leaves are scattered (usually in 2 rows) simple,
and penninerved, and with deciduous stipules.
 It is worthy of remark that in Betulaceæ, Corylaceæ and Quercus the ovules,
and to some extent the loculi of the ovary are not developed till after pollination, so
that the development of the pollen-tube proceeds very slowly. The smallness of the
flowers, the absence of honey, the dryness and lightness of the pollen, the size of
the stigma and the abundance of hairs found on many stigmas are all adaptations
for wind-pollination. It is also an advantage that the flowers are generally pollinated
before the foliage-leaves are developed, thus preventing the pollen being
entangled by the leaves.
The two orders Betulaceæ and Corylaceæ mentioned here are by other authors
united into one order. [It is doubtful whether these two should be retained in the
family Quercifloræ, as recent researches (p. 273) have shown that they differ from
the Cupuliferæ in many important points, and agree with the Casuarinas in the fact
that the pollen-tube enters the ovule through the chalaza.]

Order 1. Betulaceæ (Birches). Monœcious, with thick, cylindrical,

compound ♂ and ♀ inflorescences (2- or 3-flowered dichasia in a
spike with spirally-placed floral-leaves) (Figs. 324, 326, 328). When
the perianth in the ♂-flower is completely developed, it is composed
of 4 somewhat united leaves, which are placed opposite the 4
stamens (Figs. 325, 326 A). The female flowers are naked; the ovary
is bilocular, with two styles and one pendulous ovule in each loculus.
The subtending floral-leaves unite with the bracteoles and form a 3–
5-lobed cover-scale, which is not attached to the fruit (Figs. 325 D,
326 B). Fruit a nut without cupule (see Corylaceæ and Cupuliferæ).
In the bud the leaves are flat. The stipules are deciduous. On germination the
cotyledons are raised above the ground. Terminal buds are only found on old Alder
trees; the Birch has sympodial branches.
 Fig. 324.—Alnus glutinosus. Branch of Alder with
♂-(n) and ♀-(m) catkins: k bud; b fruit-bearing catkin
(“cone.”)
Alnus (Alder) (Figs. 324–326). In the majority of species the ♂ -
and ♀ -catkins are both developed in the year previous to their
flowering, and pass the winter naked and bloom before the leaves
expand. ♂-flower: 4 stamens. ♀-flower: the 5-lobed cover-scales of
the ♀-catkin are woody and remain attached to the axis, so that the
entire catkin when ripe resembles a small cone (Fig. 324 b). Each
cover-scale supports two winged or wingless nuts. In the native species
of Alder the buds are stalked (Fig. 324 k). The bud-scales are formed by the
stipules of the lowest leaves.
Betula (Birch). The ♂ -catkins, in the native species, appear in
autumn, the ♀ -catkins in the flowering year on leaf-bearing, short-
lived shoots. ♂-flowers: 2 stamens, divided (Fig. 328 A). The 3-lobed
cover-scales (Fig. 327 a) of the ♀-catkin are detached from the axis;
each cover-scale supports 3 broadly winged nuts (b). The stem has
cork with annual rings. The young twigs and leaves have aromatic resin glands.
 Fig. 325.—Alnus glutinosa: A dichasium of ♂-flowers seen from the front; B the
same from inside; C the same from the back; D dichasium of ♀ -flowers with
subtending-leaf and four bracteoles. The letters b, α, β, β′, β are the same as in
Fig. 326 A.
 Fig. 326.—Alnus glutinosa: diagram of dichasia of
♂ (A) and ♀ (C) catkins; B a cone-scale. All the
bracteoles in A and C are slightly pressed from their
normal position.
The Inflorescences of the Alder.—In the axil of each cover-scale [b in the
Figs] is situated, in the ♂-catkins (Figs. 326 A, 325 A-C) a 3-flowered dichasium,
the flowers of which have a 4-partite perianth, the posterior perianth-segments
being sometimes almost suppressed, and 4 stamens with undivided filaments. In
the ♀ -catkin (Figs. 325 D, 326 C) a 2-flowered dichasium is found, the middle
flower being suppressed (indicated by a star in C). In both instances the
inflorescences have two bracteoles (α-β) and the flowers borne in their axils have
each one bracteole (β′), the other one (α′) being suppressed and therefore in 326
A and C only represented by a dotted line; these four bracteoles unite with the
cover-scale (b) which supports the entire dichasium, to form the 5-lobed “cone-
scale” (Fig. 326 B) which in the ♀-catkin eventually becomes woody.
The Inflorescences of the Birch.—A 3-flowered dichasium is situated in the
axil of the cover-scale in both ♂ -and ♀ -catkins (Fig. 328 A, B); only the central
flower has bracteoles (α-β) (the lateral flowers having no bracteoles), and these
bracteoles unite, as in the Alder, with the supporting cover-scale (b), and form a
three-lobed cone-scale (Fig. 327 a).
 While the ♀-flower exactly resembles that of the Alder, the reduction of the ♂-
flower, already described in the Alder, is carried further, so that often only the 2
median perianth-leaves are developed (Fig. 328 A); there are also only 2 stamens,
these being deeply cleft, while the other 2 are suppressed.
About 50 species; N. Temp.—Fossil-forms certainly occur in the Oligocene.
During the Glacial period the Dwarf-birch (B. nana) extended over Europe; at the
present time it is confined to the moors and mountains of N. Europe and N.
America and Asia. Wind-pollinated.
Uses.—Important forest trees. The bark contains tannic acid. The tar of the
Birch is used in the preparation of Russia leather; whilst its spring sap is very
saccharine, and is used in some places for making a fermented drink. Its external
bark is used for roofing, for baskets, etc.

Fig. 327.—Betula verrucosa: a cone-scale; b fruit.